{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,4,21]],"date-time":"2026-04-21T05:15:36Z","timestamp":1776748536560,"version":"3.51.2"},"reference-count":29,"publisher":"Springer Science and Business Media LLC","issue":"7536","license":[{"start":{"date-parts":[[2014,11,12]],"date-time":"2014-11-12T00:00:00Z","timestamp":1415750400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"},{"start":{"date-parts":[[2014,11,12]],"date-time":"2014-11-12T00:00:00Z","timestamp":1415750400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springer.com\/tdm"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nature"],"published-print":{"date-parts":[[2015,1,29]]},"DOI":"10.1038\/nature13911","type":"journal-article","created":{"date-parts":[[2014,11,10]],"date-time":"2014-11-10T12:40:05Z","timestamp":1415623205000},"page":"631-634","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":188,"title":["The mitotic checkpoint complex binds a second CDC20 to inhibit active APC\/C"],"prefix":"10.1038","volume":"517","author":[{"given":"Daisuke","family":"Izawa","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Jonathon","family":"Pines","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"297","published-online":{"date-parts":[[2014,11,12]]},"reference":[{"key":"BFnature13911_CR1","doi-asserted-by":"publisher","first-page":"941","DOI":"10.1083\/jcb.130.4.941","volume":"130","author":"CL Rieder","year":"1995","unstructured":"Rieder, C. L., Cole, R. W., Khodjakov, A. & Sluder, G. The checkpoint delaying anaphase in response to chromosome monoorientation is mediated by an inhibitory signal produced by unattached kinetochores. J. Cell Biol. 130, 941\u2013948 (1995)","journal-title":"J. Cell Biol."},{"key":"BFnature13911_CR2","doi-asserted-by":"publisher","first-page":"5107","DOI":"10.1073\/pnas.94.10.5107","volume":"94","author":"CL Rieder","year":"1997","unstructured":"Rieder, C. L. et al. Mitosis in vertebrate somatic cells with two spindles: implications for the metaphase\/anaphase transition checkpoint and cleavage. Proc. Natl Acad. Sci. USA 94, 5107\u20135112 (1997)","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"BFnature13911_CR3","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1038\/10049","volume":"1","author":"P Clute","year":"1999","unstructured":"Clute, P. & Pines, J. Temporal and spatial control of cyclin B1 destruction in metaphase. Nature Cell Biol. 1, 82\u201387 (1999)","journal-title":"Nature Cell Biol."},{"key":"BFnature13911_CR4","doi-asserted-by":"publisher","first-page":"1370","DOI":"10.1038\/ncb2842","volume":"15","author":"AE Dick","year":"2013","unstructured":"Dick, A. E. & Gerlich, D. W. Kinetic framework of spindle assembly checkpoint signalling. Nature Cell Biol. 15, 1370\u20131377 (2013)","journal-title":"Nature Cell Biol."},{"key":"BFnature13911_CR5","doi-asserted-by":"publisher","first-page":"925","DOI":"10.1083\/jcb.200102093","volume":"154","author":"V Sudakin","year":"2001","unstructured":"Sudakin, V., Chan, G. K. & Yen, T. J. Checkpoint inhibition of the APC\/C in HeLa cells is mediated by a complex of BUBR1, BUB3, CDC20, and MAD2. J. Cell Biol. 154, 925\u2013936 (2001)","journal-title":"J. Cell Biol."},{"key":"BFnature13911_CR6","doi-asserted-by":"publisher","first-page":"208","DOI":"10.1038\/nature10896","volume":"484","author":"WC Chao","year":"2012","unstructured":"Chao, W. C., Kulkarni, K., Zhang, Z., Kong, E. H. & Barford, D. Structure of the mitotic checkpoint complex. Nature 484, 208\u2013213 (2012)","journal-title":"Nature"},{"key":"BFnature13911_CR7","doi-asserted-by":"publisher","first-page":"92","DOI":"10.1016\/j.molcel.2013.05.019","volume":"51","author":"JS Han","year":"2013","unstructured":"Han, J. S. et al. Catalytic assembly of the mitotic checkpoint inhibitor BubR1-Cdc20 by a Mad2-induced functional switch in Cdc20. Mol. Cell 51, 92\u2013104 (2013)","journal-title":"Mol. Cell"},{"key":"BFnature13911_CR8","doi-asserted-by":"publisher","first-page":"27","DOI":"10.1083\/jcb.201205170","volume":"199","author":"D Izawa","year":"2012","unstructured":"Izawa, D. & Pines, J. Mad2 and the APC\/C compete for the same site on Cdc20 to ensure proper chromosome segregation. J. Cell Biol. 199, 27\u201337 (2012)","journal-title":"J. Cell Biol."},{"key":"BFnature13911_CR9","doi-asserted-by":"publisher","first-page":"177","DOI":"10.1083\/jcb.201301130","volume":"201","author":"I Primorac","year":"2013","unstructured":"Primorac, I. & Musacchio, A. Panta rhei: the APC\/C at steady state. J. Cell Biol. 201, 177\u2013189 (2013)","journal-title":"J. Cell Biol."},{"key":"BFnature13911_CR10","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1038\/ncb2165","volume":"13","author":"D Izawa","year":"2011","unstructured":"Izawa, D. & Pines, J. How APC\/C-Cdc20 changes its substrate specificity in mitosis. Nature Cell Biol. 13, 223\u2013233 (2011)","journal-title":"Nature Cell Biol."},{"key":"BFnature13911_CR11","doi-asserted-by":"publisher","first-page":"227","DOI":"10.1016\/S1534-5807(01)00019-3","volume":"1","author":"Z Tang","year":"2001","unstructured":"Tang, Z., Bharadwaj, R., Li, B. & Yu, H. Mad2-independent inhibition of APCCdc20 by the mitotic checkpoint protein BubR1. Dev. Cell 1, 227\u2013237 (2001)","journal-title":"Dev. Cell"},{"key":"BFnature13911_CR12","doi-asserted-by":"publisher","first-page":"655","DOI":"10.1101\/gad.1511107","volume":"21","author":"JL Burton","year":"2007","unstructured":"Burton, J. L. & Solomon, M. J. Mad3p, a pseudosubstrate inhibitor of APCCdc20 in the spindle assembly checkpoint. Genes Dev. 21, 655\u2013667 (2007)","journal-title":"Genes Dev."},{"key":"BFnature13911_CR13","doi-asserted-by":"publisher","first-page":"e342","DOI":"10.1371\/journal.pone.0000342","volume":"2","author":"EM King","year":"2007","unstructured":"King, E. M., van der Sar, S. J. & Hardwick, K. G. Mad3 KEN boxes mediate both Cdc20 and Mad3 turnover, and are critical for the spindle checkpoint. PLoS ONE 2, e342 (2007)","journal-title":"PLoS ONE"},{"key":"BFnature13911_CR14","doi-asserted-by":"publisher","first-page":"18419","DOI":"10.1073\/pnas.1213438109","volume":"109","author":"W Tian","year":"2012","unstructured":"Tian, W. et al. Structural analysis of human Cdc20 supports multisite degron recognition by APC\/C. Proc. Natl Acad. Sci. USA 109, 18419\u201318424 (2012)","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"BFnature13911_CR15","doi-asserted-by":"publisher","first-page":"84","DOI":"10.1242\/jcs.056507","volume":"123","author":"S Elowe","year":"2010","unstructured":"Elowe, S. et al. Uncoupling of the spindle-checkpoint and chromosome-congression functions of BubR1. J. Cell Sci. 123, 84\u201394 (2010)","journal-title":"J. Cell Sci."},{"key":"BFnature13911_CR16","doi-asserted-by":"publisher","first-page":"4332","DOI":"10.1242\/jcs.094763","volume":"124","author":"P Lara-Gonzalez","year":"2011","unstructured":"Lara-Gonzalez, P., Scott, M. I., Diez, M., Sen, O. & Taylor, S. S. BubR1 blocks substrate recruitment to the APC\/C in a KEN-box-dependent manner. J. Cell Sci. 124, 4332\u20134345 (2011)","journal-title":"J. Cell Sci."},{"key":"BFnature13911_CR17","doi-asserted-by":"publisher","first-page":"1378","DOI":"10.1038\/ncb2855","volume":"15","author":"P Collin","year":"2013","unstructured":"Collin, P., Nashchekina, O., Walker, R. & Pines, J. The spindle assembly checkpoint works like a rheostat rather than a toggle switch. Nature Cell Biol. 15, 1378\u20131385 (2013)","journal-title":"Nature Cell Biol."},{"key":"BFnature13911_CR18","doi-asserted-by":"publisher","first-page":"282","DOI":"10.1074\/mcp.M700342-MCP200","volume":"7","author":"U Rothbauer","year":"2008","unstructured":"Rothbauer, U. et al. A versatile nanotrap for biochemical and functional studies with fluorescent fusion proteins. Mol. Cell. Proteomics 7, 282\u2013289 (2008)","journal-title":"Mol. Cell. Proteomics"},{"key":"BFnature13911_CR19","doi-asserted-by":"publisher","first-page":"180","DOI":"10.1016\/j.cub.2011.12.029","volume":"22","author":"DT Lau","year":"2012","unstructured":"Lau, D. T. & Murray, A. W. Mad2 and Mad3 cooperate to arrest budding yeast in mitosis. Curr. Biol. 22, 180\u2013190 (2012)","journal-title":"Curr. Biol."},{"key":"BFnature13911_CR20","doi-asserted-by":"publisher","first-page":"73","DOI":"10.1083\/jcb.201001036","volume":"190","author":"S Santaguida","year":"2010","unstructured":"Santaguida, S., Tighe, A., D'Alise, A. M., Taylor, S. S. & Musacchio, A. Dissecting the role of MPS1 in chromosome biorientation and the spindle checkpoint through the small molecule inhibitor reversine. J. Cell Biol. 190, 73\u201387 (2010)","journal-title":"J. Cell Biol."},{"key":"BFnature13911_CR21","doi-asserted-by":"publisher","first-page":"663","DOI":"10.1016\/j.devcel.2007.09.005","volume":"13","author":"T Kiyomitsu","year":"2007","unstructured":"Kiyomitsu, T., Obuse, C. & Yanagida, M. Human blinkin\/AF15q14 is required for chromosome alignment and the mitotic checkpoint through direct interaction with Bub1 and BubR1. Dev. Cell 13, 663\u2013676 (2007)","journal-title":"Dev. Cell"},{"key":"BFnature13911_CR22","doi-asserted-by":"publisher","first-page":"214","DOI":"10.1016\/j.cub.2005.01.038","volume":"15","author":"A De Antoni","year":"2005","unstructured":"De Antoni, A. et al. The Mad1\/Mad2 complex as a template for Mad2 activation in the spindle assembly checkpoint. Curr. Biol. 15, 214\u2013225 (2005)","journal-title":"Curr. Biol."},{"key":"BFnature13911_CR23","doi-asserted-by":"crossref","unstructured":"Mariani, L. et al. Role of the Mad2 dimerization interface in the spindle assembly checkpoint independent of kinetochores. Curr. Biol. (2012)","DOI":"10.1016\/j.cub.2012.08.028"},{"key":"BFnature13911_CR24","doi-asserted-by":"publisher","first-page":"387","DOI":"10.1016\/j.molcel.2004.09.031","volume":"16","author":"Z Tang","year":"2004","unstructured":"Tang, Z., Shu, H., Oncel, D., Chen, S. & Yu, H. Phosphorylation of Cdc20 by Bub1 provides a catalytic mechanism for APC\/C inhibition by the spindle checkpoint. Mol. Cell 16, 387\u2013397 (2004)","journal-title":"Mol. Cell"},{"key":"BFnature13911_CR25","doi-asserted-by":"publisher","first-page":"3905","DOI":"10.1242\/jcs.093286","volume":"124","author":"FG Westhorpe","year":"2011","unstructured":"Westhorpe, F. G., Tighe, A., Lara-Gonzalez, P. & Taylor, S. S. p31comet-mediated extraction of Mad2 from the MCC promotes efficient mitotic exit. J. Cell Sci. 124, 3905\u20133916 (2011)","journal-title":"J. Cell Sci."},{"key":"BFnature13911_CR26","doi-asserted-by":"publisher","first-page":"710","DOI":"10.1016\/j.molcel.2011.11.014","volume":"44","author":"G Varetti","year":"2011","unstructured":"Varetti, G., Guida, C., Santaguida, S., Chiroli, E. & Musacchio, A. Homeostatic control of mitotic arrest. Mol. Cell 44, 710\u2013720 (2011)","journal-title":"Mol. Cell"},{"key":"BFnature13911_CR27","doi-asserted-by":"publisher","first-page":"1234","DOI":"10.1038\/ncb2347","volume":"13","author":"J Mansfeld","year":"2011","unstructured":"Mansfeld, J., Collin, P., Collins, M. O., Choudhary, J. & Pines, J. APC15 drives the turnover of MCC-Cdc20 to make the spindle assembly checkpoint responsive to kinetochore attachment. Nature Cell Biol. 13, 1234\u20131243 (2011)","journal-title":"Nature Cell Biol."},{"key":"BFnature13911_CR28","doi-asserted-by":"publisher","first-page":"1411","DOI":"10.1038\/ncb1799","volume":"10","author":"J Nilsson","year":"2008","unstructured":"Nilsson, J., Yekezare, M., Minshull, J. & Pines, J. The APC\/C maintains the spindle assembly checkpoint by targeting Cdc20 for destruction. Nature Cell Biol. 10, 1411\u20131420 (2008)","journal-title":"Nature Cell Biol."},{"key":"BFnature13911_CR29","doi-asserted-by":"publisher","first-page":"68","DOI":"10.1016\/j.molcel.2009.02.027","volume":"34","author":"ME Matyskiela","year":"2009","unstructured":"Matyskiela, M. E. & Morgan, D. O. Analysis of activator-binding sites on the APC\/C supports a cooperative substrate-binding mechanism. Mol. Cell 34, 68\u201380 (2009)","journal-title":"Mol. Cell"}],"container-title":["Nature"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/www.nature.com\/articles\/nature13911.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/www.nature.com\/articles\/nature13911","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"http:\/\/www.nature.com\/articles\/nature13911.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,5,18]],"date-time":"2023-05-18T14:26:21Z","timestamp":1684419981000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/nature13911"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2014,11,12]]},"references-count":29,"journal-issue":{"issue":"7536","published-print":{"date-parts":[[2015,1,29]]}},"alternative-id":["BFnature13911"],"URL":"https:\/\/doi.org\/10.1038\/nature13911","relation":{"has-review":[{"id-type":"doi","id":"10.3410\/f.725230428.793501689","asserted-by":"object"}]},"ISSN":["0028-0836","1476-4687"],"issn-type":[{"value":"0028-0836","type":"print"},{"value":"1476-4687","type":"electronic"}],"subject":[],"published":{"date-parts":[[2014,11,12]]},"assertion":[{"value":"21 February 2014","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 October 2014","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"12 November 2014","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing financial interests.","order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}}]}}